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The corresponding author will normally receive a copy of the edited manuscript on which deletions, additions, and changes can be made and queries answered. Only one set of page proofs will be sent. All desired corrections of type must be made on the single set of page proofs. Changes in page proofs (as opposed to corrections) are very expensive. Author-generated changes in page proofs can only be made if the author agrees in advance to pay for them. | THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER. | GEOLOGY LIBRARY FIELDLA — as Geology wns A PAGN NEW SERIES, NO. 25 pT Ue gEaroe’ The Macropodoidea (Marsupialia) of the Early Pliocene Hamilton Local Fauna, Victoria, Australia Timothy F. Flannery Thomas H. Rich The Australian Museum Museum of Victoria 6-8 College Street 285-321 Russell Street Sydney, New South Wales Melbourne, Victoria Australia 2000 Australia 3000 William D. Turnbull Ernest L. Lundelius, Jr. Department of Geology Department of Geological Sciences Field Museum of Natural History and Vertebrate Paleontology Laboratory Chicago, Illinois 60605-2496 Texas Memorial Museum U.S.A. University of Texas at Austin Austin, Texas 78713 U.S.A. Accepted July 6, 1992 Published December 31, 1992 Publication 1443 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1992 Field Museum of Natural History ISSN 0096-2651 PRINTED IN THE UNITED STATES OF AMERICA ee ee a ee Table of Contents ABSTRACT. « 80 rte eae ies tists ts ye re ae ee eon l INTRODUCTION Sire teehee eh eg 1 MATERIALS AND METHODS) 2.05 ..3<). 22.0: c woe pu Superfamily Macropodoidea (Gray, 1821) 2 Family Potoroidae Gray, 1821 ........ 2. Subfamily Propleopinae Archer and Flannery. 1985. tess ke Z Subfamily Hypsiprymnodontinae (CEOUETE CRG Tony Sc eva sie aie See 2 Subfamily Potoroinae Trouessart, DROS oe Shere ek aor ae ety 4 Family Macropodidae Gray, 1821 ras Ot | Subfamily Sthenurinae (Glauert, 1926) 11 Subfamily Macropodinae Gray, 1821 12 TISCUSSION) Seiten ser ates sts eta ee eee 31 CONCLTISIONS 0s Saree ee Oe en cen Rin ee 34 ACKNOWLEDGMENTS: SF t ood eics cham pais 34 LITERATURE GCHED 22d on oc cen ane 34 (APPENDIX so nor Soe sate ei ainine eee ee 36 List of Illustrations 1. Hypsiprymnodon sp., SEM stereopairs OLAIPDEL IMOLATS <2 ci9 Oe Wass see ce een 3 2. Milliyowi bunganditj, stereopairs of ho- lotype:and referred specimen. . x.25.. 626. 7 3. Milliyowi bunganditj, stereopairs of teeth referred to the species ............ 8 4. Stereopairs of teeth assigned to Milli- yowi bunganditj, Troposodon sp., and SHFIOSINERUIUS SPSS eh clon tie WS oes 9 5. Drawings of teeth assigned to taxa illus- trated in Figure 4 ill 6. Dorcopsis wintercookorum, drawings of holotype and referred specimen ........ 14 7. Dorcopsis wintercookorum, stereopairs of holotype and five other specimens ... 16 8. Stereopairs of D. wintercookorum speci- mens and teeth referred cf. Dendrolagus 18 9. Thylogale ignis, stereopairs of six upper teeth, including holotype .............. 21 10. Thylogale ignis, stereopairs of lower LEC BY otc ee een rime era <p eieee 22 11. Drawings of right mandibular fragment of T. ignis and part of holotype speci- men of Kurrabi pelchenorum .......... 23 12. Kurrabi pelchenorum, drawings of holo- PYG REF Se re tae is Ga at aa ae de ee 24 13. Stereopairs of specimens referred to K. pelchenorum, crown views ............ 26 14. Drawings of teeth of K. pelchenorum BNC PTOleMNOGOK SDs ee ee ee 27 15. Drawings of teeth referred to Protemno- OPES Sn on pet eee er ee ted 30 16. Stereopairs of teeth assigned to Macro- pus (Notamacropus) sp., Wallabia sp.., and Macropodidae gen. indet. ......... 32 List of Tables 1. Dental measurements of some Hamilton PACEODOGIOS) roe Anas Reta ee 5 2. Dental measurements of Milliyvowi bun- ganditj and Aepyprymnus rufescens, com- DATO eatery scecs cee err tee eigen lavas ili! 3. Measurements of three species of Thyloga- | I, A tee aA ad get AL A ol od es a 25 4. Measurements of teeth referred to Kurra- Ui DEICREROTEIN on os state ie tn es 28 The Macropodoidea (Marsupialia) of the Early Pliocene Hamilton Local Fauna, Victoria, Australia Timothy F. Flannery William D. Turnbull Thomas H. Rich Ernest L. Lundelius, Jr. Abstract The early Pliocene Hamilton local fauna from southwestern Victoria, Australia, has at least fifteen kinds of macropodoids including propleopines, potoroines, hypsiprymnodontines, sthen- urines, and macropodines. New taxa described here include a potoroine, Milliyowi bunganditj n. gen. and sp. and the macropodines Dorcopsis wintercookorum n. sp., Thylogale ignis n. sp., and Kurrabi pelchenorum n. sp. The other ten genera present are represented by specimens for which no species assignment has been made. These are Propleopus, Hypsiprymnodon, Tropo- sodon, Simosthenurus, Dendrolagus, Protemnodon, Macropus (Notomacropus), Wallabia, and two Macropodidae gen. indet. (1 and 2). Several extant genera whose living species are almost exclusively rain forest or rain forest fringe dwellers occur in the fauna, suggesting that the fossil assemblage predominantly represents a rain forest environment. Introduction The Hamilton fossil locality was first discovered in 1952 by Edmund Gill, who found a single molar tooth. It was first identified as that of a cuscus (Gill, 1952, 1957) or phalangerid (Stirton, 1957), but it was later recognized as that of a large po- toroid (Ride, 1964). The site was subsequently worked extensively and its fauna described by Turnbull and Lundelius (1970). The fossils occur in an unnamed lithic unit that rests on the marine Grange Burn Formation and is overlain by a basalt flow. The unnamed unit is about 1.3 m thick and has been interpreted as a fossil soil (Gill, 1957). Turnbull et al. (1965) published a date of 4.35 + .1 Ma on the overlying basalt. McDougall (pers. comm., 1980) has recalculated the date as 4.46 + .1 Ma using the constants recommended for the potassium-—argon method in Steiger and Jager (1977). The geology and stratigraphy of the site were described in Turnbull and Lundelius (1970). The site is located at 37°43'58 + 03”S, 141°57'14 + 04”E, based on Sheet 7222, Coleraine 1:100,000 (Edition 1) Series R652. It is on the south side of the Grange Burn about 100 m downstream from a waterfall and approximately 7 km west of Ham- ilton, Victoria. It is indicated as site 10 on a de- tailed map (text fig. 3 in Gill, 1957). Turnbull and Lundelius (1970) also show the site in their figure 1 and identify it as the “‘Pliocene Fossil Locality.” This paper deals with the macropodoids of the Hamilton local fauna that were collected in 1966- 1967 by Turnbull and Lundelius and in 1977- 1981 by Rich and Flannery. It also includes, with some revisions, those specimens collected previ- ously and discussed in the earlier report by Turn- bull and Lundelius (1970) in which nine taxa of macropodoids were recognized. At the time only two of the nine taxa, Thylogale and Dorcopsis, could be identified with certainty to the generic level owing to the fragmentary nature of the ma- terial (mainly isolated teeth). The current collec- tion of more complete cheektooth rows in some taxa and other additional specimens has allowed the recognition of at least fourteen kinds of macro- podoids, ten of which were only identified at the generic level. While many isolated macropodoid teeth are present in the collections, recognition of FIELDIANA: GEOLOGY, N.S., NO. 25, DECEMBER 31, 1992, PP. 1-37 1 their proper taxonomic assignments is generally uncertain because the teeth of the taxa are very similar. Hence, only distinctive or associated teeth are discussed extensively in this paper. The many isolated teeth of unsure affinity are listed in the Appendix. Only five Pliocene faunas with a moderately di- verse macropodoid assemblage have been report- ed from Australia-New Guinea. These are the Hamilton local fauna from Victoria, the Bow local fauna from New South Wales (Skilbeck, 1980; Flannery & Archer, 1984), the Chinchilla and Bluff downs local faunas from Queensland (Bartholo- mai, 1963, 1967, 1973, 1975, 1976, 1978b), and the Awe local fauna from New Guinea (Plane, 1967). Hamilton is the most firmly dated locality and contains the second greatest diversity of mac- ropodoids (after Bow), thus making it one of the most informative of the sites from the viewpoint of macropodoid evolution. The Hamilton site has produced almost exclusively mammalian and plant remains, the exceptions being several frog bones (which may possibly be modern). Bone was poorly preserved at the site, the most common remains being tooth caps. Mammalian groups represented are also diverse: Included are the Dasyuridae, Peramelidae, Petauridae, Burramyidae, Phalan- geridae, Ektopodontidae, Vombatidae, Potoro- idae, Macropodidae, Palorchestidae, Diprotodon- tidae, and Microchiroptera. Materials and Methods All specimens of macropodoids from the Ham- ilton local fauna not cited in the text are listed in the Appendix. Abbreviations are PM— fossil mam- mal collection, Field Museum of Natural History, Chicago, and NMv P—paleontological collection, Museum of Victoria, Melbourne. The diagnoses given below are not meant to be exhaustive, but cover only the fossil elements being diagnosed. Enumeration of cheek teeth differs from that in Turnbull and Lundelius (1970) and follows Archer (1978). The posterolingual cusp of the upper mo- lars is here designated the hypocone, pending fur- ther work concerning the suggestion of Tedford and Woodburne (1987) that it is the metaconule. Paracone and metacone are retained for the buccal cusps of the protoloph and metaloph, although they may ultimately prove to be stylar cusps (Ted- ford & Woodburne, 1987). Measurements were taken with dial calipers. Superfamily Macropodoidea (Gray, 1821) Family Potoroidae Gray, 1821 Subfamily Propleopinae Archer and Flannery, 1985 Propleopus Longman, 1924 Propleopus sp. DISCUSSION—A_ species of Propleopus repre- sented by isolated molars and premolars from the Hamilton local fauna has been under study by W. D. L. Ride and the late J. A. Mahoney. The first tooth (NMv P15777) discovered at Hamilton, which was recognized by Ride (1964) as the molar of a large potoroid, probably belongs within this taxon. Eight other specimens (PM 16817, PM 16819, PM 16820, PM 16832, NMv P150212, NMv P150213, NMV P150274, and NMv P150296) have been sent to Ride for incorporation into that study, which will appear elsewhere. Subfamily Hypsiprymnodontinae (Collett, 1877) Hypsiprymnodon Ramsay, 1876 Hypsiprymnodon sp. MATERIAL— PM 4575, protoloph of right M?; NMV P169186, right M, (fig. 1A); NMv P160153, pro- tolophid of right M, or protoloph of M?; pm 50762, nearly complete left upper molar (fig. 1B). DESCRIPTION—A large, distinct trigonid basin is present on NMV P169186, the M,, and the anterior cingulum is rounded in occlusal view. There is no fossette labial to the paracristid. The posterior cin- gulum is small and encloses a small posterior fos- sette. The protolophid is longer than the hypolo- phid, which is barely discernable at its midpoint. The four major cusps are located more to the out- side of the tooth than is the case in other poto- roines. The relative sizes of the four principal cusps are protoconid > metaconid > hypoconid > en- toconid. Ride (1961) implies that the protoconid and metaconid are of equal size in the living Hyp- siprymnodon moschatus. The cristids from the protoconid and hypoconid join to form a low wall on the buccal side of the talonid basin. The cristids from the metaconid and the entoconid do not meet on the lingual side of the talonid basin. The upper molar (PM 50762) has a protoloph that is longer than the metaloph, which is barely discernable at its midpoint in the median valley. Part of the metacone is missing, but the tooth is FIELDIANA: GEOLOGY Imm Fic. 1. Hypsiprymnodon sp. Scanning electron microscope stereopairs. A, NMV P160186, right M‘ shown in crown view at x25. B, pm 50762, left upper molar shown in crown view at x29. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 6) nearly complete. There is a broad precingular ba- sin, a large central basin between the lophs, and a distinct basin posterior to the metaloph. Each loph is interrupted by a fissure that crosses the loph at its lowest point. There is a small labial basin be- tween the paracone and metacone. Labiad, the weak crests that run from the apices of these cusps meet in the transverse valley. This labial basin lies just labiad to the junction of the crests and linguad to the labial cingulum. A groove on the lingual side of the tooth, between protocone and hypocone (= metaconule of Tedford & Woodburne, 1987), serves to extend the transverse valley onto the lingual side of the crown, but the crests from those cusps interrupt the valley and groove where they join. Cusps and lophs are low, and the enamel in the valleys shows some crenulation. DIscUSSION— These molar fragments and mo- lars have been referred to Hypsiprymnodon sp. because they are distinctly bunodont in form and are smaller than those of any potoroine or species of Propleopus and the lophs/ids are less well formed. This combination of characteristics is unique to Hypsiprymnodon moschatus in the extant Macro- podoidea. Furthermore, the M* is more elongate than in species of Bettongia, Aepyprymnus, Cal- oprymnus, Milliyowi n. gen., or Propleopus. Al- though the upper and lower molars and molar fragments are bunodont in form, the cristae orig- inating from the hypocone/id and protocone/id are poorly developed, especially when compared to those in species of Caloprymnus, Propleopus, Aepyprymnus, Wakiewakie, Palaeopotorous, Bu- lungamaya, and Milliyowi. The almost quadritu- bercular molar pattern resulting is only matched among macropodoids in H. moschatus. A taxon similar to H. moschatus was recognized by Turnbull and Lundelius (1970) on the basis of PM 4575, the protoloph of an M?. The additional material described here has allowed a more certain identification as Hypsiprymnodon sp. While this material is easily distinguishable from most po- toroines on the basis of morphology, it is primarily the smaller size and weaker loph/id development that distinguishes it from species of Potorous (see table 1). Subfamily Potoroinae Trouessart, 1898 Milliyowi n. gen. Type Species— Milliyowi bunganditj n. sp. GENERIC DIAGNOsIS— Milliyowi bunganditj can be distinguished from all other potoroines in pos- sessing anterior upper molars with extended slop- ing lingual edges and anterior lower molars with sloping labial edges. It differs from the species of Potorous and Caloprymnus (which have four to five poorly defined ridges) in having at least seven well-defined ridges on P°. It differs from species of Bettongia except (variably) B. gaimardi in lack- ing a posterolingual cuspule on P?. The teeth of M. bunganditj are closest in size to those of Ae- pyprymnus rufescens among Potoroinae but are slightly larger (table 2). While most similar in over- all morphology to A. rufescens, the molars of M. bunganditj differ, in addition to the above char- acters, in having a more gently sloping posthy- pocristid and posthypocrista. The posterior cin- gulum, the metaconule, and the hypocone/id are less massive and the teeth have more open and more centrally located cingular basins than in A. rufescens and, to a lesser extent, C. campestris. It differs from Gumardee pascuali in having a P? that lacks buccal and lingual cingulae and a large bulge on the posterolingual corner (Flannery et al., 1982). The P? differs from that of Wakiewakie lawsoni in the lack of a labial turn at the posterior end, in having an incipient posterolabial cusp, and in that the ribbing is coarser. Milliyowi differs from Palaeopotorous priscus in the larger size of the molars, in the absence of a postlink, a midlink, and a protoconule on the upper molars, and in that M, lacks any sign of a protostylid. ETYMOLOGY —From the aboriginal millia (rat- kangaroo) and yowi (spirit surviving the death of the body) (Parker, 1953). Milliyowi bunganditj n. sp. (figs. 2, 3, 4A, SA; table 2) HOLOTYPE—NMV P158635, associated in a sin- gle lump of matrix are enamel caps of the posterior two-thirds of right P?, and right M3 (figs. 2A,C,E). REFERRED SPECIMENS— PM 50763, right M! (figs. 2B,D); NMv P158068, right M? (fig. 3A); Pm 16821, crown of a left I, (fig. SA); Nuv P53223, anterior three-quarters of right P, (figs. 3B—D); NMv P158069, right M, lacking anterior portion of pro- toconid (fig. 3E); Nuv P54137, left M,; or M, (fig. 4A). TYPE LOCALITY AND AGE—See the Introduction for details. DIAGNosis— The diagnosis of the genus will serve as that of the species until further species are rec- ognized. FIELDIANA: GEOLOGY TABLE 1. Dental measurements (mm) of some Hamilton macropodids. L AW PW Hypsiprymnodon sp. NMV P160186 M, 2.8 1.9 22 cf. Dendrolagus NMV P157483 Fr? 4.9 NMV P157481 BP 4.2 Dorcopsis wintercookorum PM 16822 ee 9.8 3.4 3.6 NMV P158453 Pp 9.8 3.3 4.0 2 4.4 4.2 4.0 M3 4.7 4.3 3.9 Mé 4.8 4.6 4.0 MS 4.9 4.1 3:3 PM 50771 M* 5:2 4.5 3.8 NMV P157477 P; 9.5 2.4 3:2 NMV P158462 RP 8.5 22 25 NMV P160221 M'! 4.8 3.6 3.8 PM 16816 M'! 5.0 3.6 3.9 PM 50767 2 4.8 4.0 4.3 Troposodon sp. NMV P158465 M> 9.8 Simosthenurus sp. NMV P158633 Hypolophid 8.6 NMV P158594 Protolophid 9.6 Protemnodon sp. NMV P160372 M'! 9.9 10.1 8.5 Macropus (Notamacropus) sp. NMV P54112 Hypolophid da3 cf. Wallabia NMV P54208 ip? 8.6 S33 4.2 Gen. indet. NMV P158431 Pe 4.4 1.8 Mey L = length, AW = anterior width, PW = posterior width. ETyMOLOGY — Bunganditj, the name of the ab- original tribe that used to inhabit the Hamilton area (West, 1974). DESCRIPTION— P? — The posterior portion of the P? (fig. 2A) present in the holotype is ornamented with seven well-developed ridges labially and there is a suggestion of an eighth lingually at the broken surface in front. By comparison with A. rufescens, it appears that the fragment represents the pos- terior two-thirds of the tooth. The P® of A. rufes- cens has seven or eight ridges. It is certain that the complete P* of M. bunganditj had at least one more ridge than is preserved, and it perhaps had as many as ten if we are correct in our interpretation of the portion represented. The P*® is weakly concave buccally and lingually and is much narrower buc- colingually than that of species of Propleopus. The ridges form cuspules where they meet on the straight occlusal crest, except posterolingually. No posterolingual cusp is present, but the tooth thick- ens toward its posterior edge. The presence of a posterolingual cusp on the P® is variable in A. ru- fescens and B. gaimardi, and this character also may have been variable in M. bunganditj. Table 2 gives comparative measurements for the den- titions of species of Milliyowi and Aepyprymnus. Upper Molars—The M! (figs. 2B,D) is an un- worn, fully formed crown, clearly from a very young individual. The parastyle of M' is very elongate and massive. There is no suggestion of an anterior cingulum, and the anterior face of the protoloph slopes steeply to the crown base. The protoloph is low and poorly developed, has a small, posteriorly directed kink, and is shorter than the well-devel- FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 5 oped metaloph. A very prominent postparacrista is prevented from linking with a more weakly de- veloped premetacrista by a deep fissure. The mid- link is well developed, as is the metaloph. The lophs are noticeably narrow at their apices. The ectoloph is interrupted by a narrow valley that connects the interloph valley to the labial side of the crown. Anteriorly, the ectoloph divides near its end. The metaloph is noticeably kinked and the protocone is small, being much the smallest of the primary cusps. The rectangular central val- ley is longer than wide. A broad stout postcingu- lum defines a broad arcuate posterior basin. A slightly developed postlink is present on the rear face of the metaloph. The M? (fig. 2C) is barely worn. The primary lophs are subequal in length and the protoloph is well formed but its connection to the base of the protocone is weakly developed. The pre- and post- protocristae are well developed. The preproto- crista forms a well-developed anterior cingulum that unites with a weakly developed preparacrista on the labial margin of the tooth. The postpara- crista and premetacrista are weakly developed. They do not join at the base of the tooth, leaving the central basin open buccally. In contrast, the pre- and posthypocristae are well developed, the posthypocrista forming a posterior cingulum by uniting with the postmetacrista. The prehypocrista and postprotocrista join close to the lingual side of the central basin. The metaloph is well devel- oped. The lingual margin of the tooth slopes gently to the crown base. The M3? (fig. 2E) is less worn than the M? and differs from it in having a more steeply sloping lingual margin. A fissure can be seen in the pro- toloph and another in the metaloph, both near their lingual edges. To judge by the associated val- leys, this feature was probably present on M? be- fore becoming obscured by wear. The M® (fig. 3A), if properly assigned to this taxon, differs from M? in having a steeper lingual margin and in being lower-crowned. The metaloph is shorter than the protoloph, and the ridges and lophs are sharper and less bulbous than in the other specimens. I, —The extremely elongate and narrow enamel cap of an I, (fig. SA) appears to belong to a potoroid approximately the size of Milliyowi bunganditj and is here tentatively referred to that species. The enamel cap is most like that of I, of a species of Potorous. It differs from I, of the species of Wakie- wakie, Bettongia, Caloprymnus, and Aepyprymnus in possessing a slight dorsal enamel crest extending the length of the crown as in some macropodids. It differs from I, of all macropodids by being nar- rower and more elongate. The I, of Gumardee pas- cuali is unknown, while a poorly preserved I, has been reported for Bulungamaya delicota (Flannery et al., 1982). It is clear, however, that the I, of B. delicota is less elongate than that of M. bunganditj. P,—The P; (figs. 3B—D) is represented by a frag- ment that consists of the anterior three-fourths of a tooth. It has a nearly flat straight blade with six sharply defined cusplets and nearly vertically ori- ented associated ridges. The remnant of a seventh ridge can be seen on the labial side. Small subsid- iary branches come off of most of the main ridges as they approach the crest. These disappear in the intervening valleys. This branching feature is not seen in any of the comparative specimens of A. rufescens, Wakiewakie, Potorous, Bettongia, Bu- lungamaya, or Caloprymnus available to us. A sharp ridge defines the anterior edge of the tooth. The blade is buccolingually narrow. Lower Molars— The anterior cingulum and buc- cal portion of the protolophid of the M, (NMv P158069) are broken away (fig. 3A). The labial face of the hypolophid slopes at a gentle angle to the base of the crown. The protolophid appears to — Fic. 2. Milliyowi bunganditj. Stereographic photopairs. All approx. x6.8. A, C, E, Holotype specimen, NMv P158635, associated right P?, M?, and M? shown in crown view. B, D, Referred specimen, PM 50763, a right M'! shown in crown and buccal views. Fic. 3. Milliyowi bunganditj. Stereographic photopairs of three teeth. C is shown at approx. <6.7, the rest at x 8.3. A, NMV P158068, a tentatively referred right M> shown in crown view. B-D, NMv P53223, right P; shown in buccal, lingual, and crown views. E, NMv P158069, a posterior two-thirds of a right M,, lacking anterior part of protoconid, shown in crown view. (overleaf) Fic. 4. Stereographic photopairs of teeth of Milliyowi bunganditj (A), Troposodon sp. (B, C), and Simosthenurus sp. (D, E), all shown in crown view at approx. <6.7. A, NMv P54137, left M,. B, NMv P158465, right M,. C, NMV P26422, hypolophid of a left lower molar. D, NMv P158449, a protolophid of a left lower molar. E, NMv P158633, hypolophid of a right lower molar. (page 9) FIELDIANA: GEOLOGY 3mm FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE qi ! } “ = ate tt ec nt, ‘a FIELDIANA: GEOLOGY FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 1cm Fic. 5. Drawings of various teeth representing the same three genera as in Figure 4. All approx. x6.7. A, PM 16821, a left I, of Milliyowi bunganditj shown in ventrolateral and dorsomedial views. B, NMv P158465, the same specimen as Figure 4B, right M* of Troposodon sp. shown in lingual, crown, and posterior views. C, NMv P158449, the same specimen as in Figure 4D, protolophid of a left lower molar of Simosthenurus sp. shown in crown and anterior views. D, NMv P158633, the same tooth as in Figure 4E, hypolophid of a right lower molar of Simosthenurus sp. shown in crown and posterior views. 10 FIELDIANA: GEOLOGY : ‘ 3 s TABLE 2. Dental measurements (mm) of Milliyowi bunganditj compared to those of Aepyprymnus rufescens. Aepyprymnus rufescens Milliyowi bunganditj >, ¢ R SD N NMV P158635 Pe PW 4.0 3:2 2.9-3.6 23 7 M? if 6.0 55) 5.1-6.0 soil 12 AW 6.4 4.3 4.0-4.5 14 12 PW 6.7 4.1 3.9-4.5 18 12 M? L 6.8 6.3 5.9-6.6 .22 12 AW ae 4.7 4.4-5.0 21 12 PW 5.6 4.3 4.0-4.3 22 12 NMV P158068 M> L 6.5 6.2 5.7-6.6 — 2 AW a3 4.2 4.1-4.3 = 2 PW 4.2 3.3 3.1-3.5 _ 2 PM 50763 M'! L 5.4 = _ _ _ AW 4.0 _ = = _ PW 4.5 3.3 3.1-3.5 2 3 NMV P158069 M! i b:>:5.6 AW b> 74.1 PW a1 NMV P54137 M, L 6.7 6.3 6.0-6.9 a32. 8 AW 4.9 4.8 4.4-5.2 .26 7 PW 5:0 4.5 4.2-5.0 .26 8 L = length, AW = anterior width, PW = posterior width, K = mean, R = range, SD = standard deviation, N = number, b = broken. form a single cusp as in other potoroines. The cristid obliqua terminates before reaching the pro- toconid apex. The hypolophid is poorly devel- oped. A slight postprotocristid and a preentocris- tid are present. The presumed M, or M, (fig. 4A) has a buccal edge that slopes at a steeper angle to the crown base than in M,. The paracristid forms the labial and anterior boundary of the extensive anterior cingulum. There is no fossette buccal to this struc- ture as found in macropodids and some other po- toroids (e.g., C. campestris, bulungamayines). The postprotocristid is well developed and joins a well- developed prehypocristid to form the cristid ob- liqua. The posthypocristid forms a well-developed posterior cingulum. A slight low point can be seen on both protolophid and hypolophid near their labial margins. The central valley has ornamen- tation on the lingual side. DiscussioN—The relationships of M. bungan- ditj are at present unclear. Flannery (1989) in- cluded Milliyowi bunganditj within his analysis of the phylogeny of the Macropodoidea, giving it the informal name the “Hamilton potoroid.” He found it difficult to place because of the large number of features that could not be examined due to the fragmentary nature of the fossils. He did note, however, that it lacks a significant synapomorphy FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE that unites Aepyprymnus and Caloprymnus. This is hypertrophy of the posthypocristid on the lower molars, which in these genera slopes diagonally across almost the entire width of the hypolophid. Thus Milliyowi appears to lie outside the Aepy- prymnus/Caloprymnus clade. We tentatively place it within the tribe Bettongini, but leave it incertae sedis within that group. A single upper molar frag- ment (PM 4492), referred to as cf. Aepyprymnus by Turnbull and Lundelius (1970), may be refer- able to M. bunganditj; however, it is too fragmen- tary for any assignment to be certain. Family Macropodidae Gray, 1821 Subfamily Sthenurinae (Glauert, 1926) Troposodon Bartholomai, 1967 Troposodon sp. MATERIAL—NMV P158465, right M* missing the anterior side of the protoloph and much of the protocone (figs. 4B, 5B); NMv P26422, posterior half of a right lower molar (fig. 4C; ref. Turnbull & Lundelius, 1970, p. 63). DESCRIPTION— The more complete molar (figs. 4B, 5B) lacks the anterior face of the protoloph and the protocone and the surrounding area. Enough is preserved to show that the lophs are 11 attenuated, the midlink is well developed, and a well-developed postprotocrista is present. The posthypocrista and postmetacrista overlap and a prominent postlink is present in the center of the rear face of the hypoloph. The postparacrista ex- tends from the crest of the paracone posterolin- gually on the posterior face of the protoloph and disappears before reaching the midlink. The tooth is much larger than molars of Lagostrophus fas- ciatus, and it differs from upper molars of species of Simosthenurus, Procoptodon, and to some ex- tent Sthenurus in lacking the fine ornamentation and/or grooving of the enamel seen in these forms. There is barely discernable ornamentation on the posterior face of the protoloph. DiscussiON—Turnbull and Lundelius (1970) suggested the presence of a species of Troposodon in the Hamilton local fauna on the basis of a hy- polophid (NMv P26422). While the two molar frag- ments discussed here are close in morphology to Troposodon minor, they are too incomplete to per- mit assignment to a species. The combination of features seen in the upper molar fragment, includ- ing its large size, overlapping of a distinct posthy- pocrista and postmetacrista, very prominent post- link, and lack of any enamel crenulation, is unique to the species of Troposodon among Macropo- doidea. Simosthenurus Tedford, 1966 Simosthenurus sp. (figs. 4D,E, 5C,D; table 1) MATERIAL—NMV P158449, left protolophid (figs. 4D, 5C); NMv P158633, right hypolophid (figs. 4E, SPD): DESCRIPTION—The two molar fragments are low-crowned; the protolophid extremely so. Their features most clearly resemble those of Simos- thenurus antiquus but they are smaller. Other sim- ilarities are the posterior position of the preme- tacristid and the distinct paracristid of NMV P158449, and the sharp cristid obliqua of NMV P158633. The trigonid basin is broad, and both protolophid and hypolophid are ornamented with fine crenulations. The paracristid originates well below and lingual to the protoconid apex. The premetacristid crosses the face of the protolophid, a primitive characteristic. The hypolophid is bul- bous at its base. DISCUSSION — This material has been assigned to a species of Simosthenurus because the enamel of the lower molar fragments is finely crenulate (a LZ feature seen among macropodids only in the spe- cies of Procoptodon and Simosthenurus) and be- cause the posterior face of the hypolophid lacks the deep grooving seen in the species of Procop- todon (Flannery, 1983). Also, the molar fragments are lower-crowned than those of species of either Sthenurus or Procoptodon. However, it differs from all other species of Simosthenurus (as well as Pro- coptodon and Sthenurus) in having the premeta- cristid located on the anterior face of, and oriented at about 45° to, the protolophid. Simosthenurus antiquus is the only named Plio- cene species of the genus. However, isolated molar and premolar fragments of an unnamed species of Simosthenurus are known from the Bow local fau- na (Flannery & Archer, 1984) as well as from Hamilton. The Hamilton material is more prim- itive in its low-crowned condition than other forms and represents a new species, but more complete material is needed before it can be named. The Hamilton molar fragments represent the earliest known occurrence of a species of Simosthenurus. Subfamily Macropodinae Gray, 1821 Dorcopsis Schlegel and Miller, 1845 Dorcopsis wintercookorum n. sp. (figs. 6-8; table 1) HOLOTYPE—NMV P158453, right maxilla with P>, M?> (figs. 6A—C, 7B,C). The maxilla is badly corroded, but the enamel caps of the teeth are in good condition. REFERRED SPECIMENS—PM 16822, left P® (figs. 6D-F, 7A); NMv P160221, left M! (fig. 7E); pm 16816, right M! (fig. 7D); pM 4570, protoloph of right M?; pm 50767, right M? (fig. 7F); pm 16807, protoloph of right upper molar, probably M? or M3; PM 50771, right upper molar (fig. 7G); NMv P158462, right P; (figs. 8B—D); NMv P157465, an- terior portion of right P,;; NMv P157477, right P; (fig. 8A). TYPE LOCALITY AND AGE—See the Introduction for details. DIAGNOsIS— Dorcopsis wintercookorum can be distinguished from other species of Dorcopsis in possessing the following characteristics. It is much smaller than any other species and possesses a cuspule labial to the paracone on M? that is not seen in any other form. The anterior edges of the P¥, of D. wintercookorum slope from the base of the crown to the occlusal margin more gently than in other species. D. wintercookorum possesses a much less distinct lingual cingulum on the P? than FIELDIANA: GEOLOGY does D. mulleri. D. hageni is closest in morphology to D. wintercookorum but is much larger (see table 1 for dental measurements of D. wintercookorum). Features that distinguish Dorcopsis wintercook- orum from Dorcopsulus are few. It has slightly larger upper molars, which are otherwise nearly identical in form to those of Dorcopsulus. The cus- pule labial to the paracone of the M? of Dorcopsis wintercookorum, which is distinctive for that tax- on alone among the species of Dorcopsis, is even better developed in the M?’s of Dorcopsulus. In that genus the cuspule is well developed on M?°’s and is present to well developed on the M*’s, and in one specimen in the American Museum of Nat- ural History sample of seven, it is present on the M?’s (table 1). In the P3’s of Dorcopsis wintercook- orum, the main crest has a variable number of cuspules (five or six). In Dorcopsulus there appear to be six. ETYMOLOGY—Named for Mr. and Mrs. Sam Wintercook, whose hospitality made our field- work much more enjoyable in the years from 1977 to 1981. DESCRIPTION— Maxilla— Although the maxilla | of the type specimen is badly corroded, some fea- tures can be recognized. The zygomatic process appears to be short and rounded as in the living species of Dorcopsis, not reaching the base of the molar crowns. Sutures, the position of the infra- orbital canal, and details of the palatal area are all obscured. P?— The P® of D. wintercookorum consists of a blade with a distinct, sharp anterior cusp and a more elongate posterior cusp, each with labial and lingual ridges (figs. 6A—F, 7A—C). Three distinct but weak ridges are present on the sides of the blade, and the associated cuspules cap the crest of the blade. Several low tubercles of enamel are pres- ent on the labial side of the base of the crown. A posterolingual cusp, which is lower than the main crest, is also present. It is connected to the main crest by a single ridge, which intersects the main crest at a 90° angle. The lingual cingulum is very weakly developed. It terminates anteriorly just posterolingual to the main cusp. The isolated P* is similar to that of the holotype but its lateral ridges are somewhat sharper. Upper Molars—The relative positions of the molars of the holotype are slightly distorted. How- ever, the cheektooth rows appear to have been straight and were probably parallel. Except for the M! the descriptions are based largely on the ho- lotype. The two M! specimens (figs. 7D,E) are low- crowned. A distinct and large parastyle is present. The protoloph is weakly developed, the paracone and protocone forming distinct prominences. Well- developed postparacrista and premetacrista block the lingual end of the interloph valley. The midlink is extremely poorly developed, but the metaloph is well developed. The postmetacrista and posthy- pocrista form a “U’’-shaped posterior cingulum. A postlink is present on the rear face of the me- taloph midway between the metacone and meta- conule. The two halves of the tooth are nearly equal in NMV P160221, but in pm 16816 the an- terior is slightly narrower than the posterior. The M? differs from M!' in the following ways. The M? of the holotype, which is considerably worn, is slightly larger than the M! described above. The protoloph is well developed, the parastyle is greatly reduced in size, and the midlink is slightly better developed. A small forelink is present, and posterobuccal to the paracone there is a small cus- pule that is approximately one-half the height of the paracone. The postparacrista and premeta- crista are smaller in size. Both M? specimens pos- sess a stout cingular cuspule labial to the paracone. The M3? differs from M? in the following ways. It is very slightly larger. The anterior cingulum in that of the holotype is broken away. There is no cuspule buccal to the paracone. The postparacrista and premetacrista are slightly smaller than in M?. The M‘* is similar in morphology to M°, differing only in that the postparacrista and premetacrista are further reduced in size. The M? differs from M* in that the metaloph is markedly narrower than the protoloph, and in that the postparacrista and premetacrista are still fur- ther reduced. P,—Two complete and one partial P; of Dor- copsis wintercookorum are known. Both complete specimens have a sharp and distinct anterior cus- pid and a longer posterior cuspid at the end of the main blade (figs. 8A—D). The anterior and poste- rior cuspids have distinct lingual and labial ridges from tip to cingulum. Three indistinct ridgelets are present between these cuspids (and there is a suggestion of one more in one specimen). Ante- riorly the blade slopes at a low angle to the base of the crown. The posterior portion of the tooth is slightly broader than the anterior portion. A small, low posterolabial cuspid is present on NMV P154477. The blades of both teeth are slightly convex buccally. DiscUSSION—The material assigned here to D. wintercookorum is significant in several ways. Bar- tholomai (1978a) suggested that Dorcopsoides FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 13 11cm (& eal | Fic. 6. A-C, Drawings of the holotype specimen of Dorcopsis wintercookorum, NMV P158453, a right maxillary fragment with P?, M2-> shown in buccal (A), crown (B), and lingual (C) views at approx. x 6.3. D-F, Drawings of a referred specimen of D. wintercookorum, PM 16822, shown in lingual (D), crown (E), and buccal (F) views at approx. x 8.6. 14 FIELDIANA: GEOLOGY 1cm Fic. 6. Continued. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE is 5mm Icm 5mm Fic. 7. Stereophoto pairs of various specimens of D. wintercookorum, all in crown view. A, PM 16822, left P® shown at approx. x 4.1. B, C, NMv P158453, the holotype specimen. In B, the P? is shown at the same scale as A to 16 FIELDIANA: GEOLOGY should be placed in synonymy with Wallabia, thus challenging the only previous evidence that mem- bers of the Dorcopsis lineage have inhabited Aus- tralia. He suggested that study of the Hamilton material assigned to Dorcopsis might provide clar- ification as to whether or not the Dorcopsis lineage has ever been present in Australia. The Hamilton material does indeed provide insights into this question. It is also significant in providing insights into the taxonomy of the living genera Dorcopsis and Dorcopsulus. We have no doubt that Dorcopsis wintercooko- rum isa member of the Dorcopsis lineage and bears no close resemblance to species of Wallabia. In his synonomy of Dorcopsoides with Wallabia, Bar- tholomai noted the following features in Dorcop- soides, which he considered indicated an affinity with Wallabia: the permanent cheek teeth of the two genera are very similar, the length of the pre- molar relative to the molars is more like that in Wallabia, not Dorcopsis, the molars are more like those of Wallabia, not Dorcopsis, the molars of Dorcopsoides are larger than those of Dorcopsis, Dorcopsoides differs from Dorcopsulus in that the molars vary greatly in size along the molar now, the mandible is deep and the diastema short as in Wallabia, the molar row is bowed labially, and what is known of the cranium is similar to Wal- labia. In all of these features where they can be observed, Dorcopsis wintercookorum is similar to the Dorcopsis lineage and differs from the condi- tion seen in Wallabia. The permanent cheek teeth are very similar to those of Dorcopsis and Dor- copsulus; the molars vary little in size, are very low-crowned, and lack the extreme development of linking along the labial edge of the crown seen in Wallabia. The premolars are similar in size to Dorcopsis relative to the molars and longer than in Wallabia. The upper premolar is relatively lon- ger and narrower than in Wallabia and has a very weakly developed lingual cingulum, again similar to the condition seen in Dorcopsis, but differing from the prominent and raised lingual cingulum of the species of Wallabia. The teeth are very small, being similar in size to those of Dorcopsulus van- heurni and much smaller than those of Wallabia, and the cheektooth rows are parallel, not bowed as in Wallabia. Unfortunately the dentary and cra- nium of D. wintercookorum are not known. How- ever, the evidence from cheektooth morphology seems conclusive: D. wintercookorum is a member of the Dorcopsis lineage and, regardless of the af- finities of Dorcopsoides, it provides clear evidence that this lineage once existed in Australia. Questions regarding the taxonomy of members of the Dorcopsis lineage are not so easily resolved. In his revision of the Macropodidae, Tate (1948) indicated that Dorcopsulus and Dorcopsis are very close in morphology, differing primarily in that the P? of Dorcopsulus is less compressed and at- tenuated than in Dorcopsis, and the skull is less elongate. Examination of a large series of Dorcop- sis and Dorcopsulus held at the Australian Mu- seum, Sydney, shows that premolar shape varies considerably within both genera and that few fea- tures except size reliably serve to distinguish living members of these genera. In general, however, the labial ridging of P* is more prominent in Dorcop- sulus, and M?~ (and occasionally M°) have a cus- pule positioned posterobuccal to the paracone, a feature generally lacking in Dorcopsis. Indeed, the features that serve to distinguish these genera are small, and it may be that future taxonomic revi- sion will see the genera synonymized, in which case Dorcopsis Schlegel and Miller, 1845, would take priority over Dorcopsulus Matschie, 1916. The morphology of Dorcopsis wintercookorum further muddies the distinction between these two genera. It is the size of Dorcopsulus, yet the per- manent premolars have relatively poorly defined buccal ridging, typical of Dorcopsis, and differing from Dorcopsulus, which has more prominent ridges. Of the upper molars, only M? has a cuspule positioned posterobuccal to the paracone, and even that cuspule is small relative to that seen in Dor- copsulus. In view of the uncertainty surrounding the distinctness of the two genera, and the fact that Dorcopsis is the prior name, we have assigned D. wintercookorum to Dorcopsis pending a full revi- sion of the two genera. Dorcopsis sp. REFERRED SPECIMENS—PM 4436, right P® (fig- ured by Turnbull & Lundelius, 1970, pl. XIX); NMV P26417, right upper molar (Turnbull & Lun- delius, 1970, pl. XXI). Both are larger than D. — facilitate direct comparison. In C, the series P?, M?-° are shown at approx. x 2.3. D, pM 16816, right M' shown at approx. x 4.1. E, NMv P160221, left M' shown at approx. x 6.8. F, pm 50767, right M? shown at approx. x 6.8. G, PM 50771, right upper molar (position uncertain) shown at approx. 4.1. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 17 Fic.8. Stereophoto pairs of various specimens of D. wintercookorum (A-D) and specimens referred cf. Dendrolagus (E, F). A, NMv P157477, right P, shown in crown view at approx. x 4.2. B-D, NMv P158462, right P; shown in crown 18 FIELDIANA: GEOLOGY wintercookorum, and the P® has a different mor- phology. DiscussIoN—Turnbull and Lundelius (1970) recognized the presence of Dorcopsis in the Ham- ilton local fauna on the basis of a nearly complete lower molar and two fragmentary lower molars, three complete and nine fragmentary upper mo- lars, and a fragmentary P*. The upper molars de- scribed by Turnbull and Lundelius are similar to those of the holotype of D. wintercookorum, but no comparison with the lower molars can be made. The P? (pm 4436) differs markedly from those de- scribed here for D. wintercookorum and in some ways resembles the P*’s referred below to cf. Den- drolagus sp. Turnbull and Lundelius (1970) orig- inally referred PM 4436 to Dorcopsis sp. This is continued here. This specimen differs from the P* of the other specimens of Dorcopsis wintercook- orum in possessing a broad, bladelike postero- lingual cuspid that approaches the main crest at a 60° angle (not 90° as in D. wintercookorum) and in its broad lingual cingulum. Except for the angle, in these characteristics it resembles the premolar fragments here assigned to cf. Dendrolagus sp., but it shows no sign of a labial (buccal) cingulum or, perhaps more significantly, the bladelike labial crest that Dendrolagus usually shows. The existing sam- ple of these Dendrolagus-like specimens is too fragmentary to permit adequate description of a new species. Dorcopsis wintercookorum and Dorcopsis sp. are the only species of Dorcopsis known from Austra- lia, all living species and other possible fossil oc- currences (Woodburne, 1967) being restricted to New Guinea. The presence of Dorcopsis in south- ern Australia during the early Pliocene suggests that species of the genus were at that time wide- spread and that they have subsequently become extinct in all but the northern, more tropical end of their former range, probably in response to cli- matic change. Dendrolagus Miller and Schlegel, 1839 cf. Dendrolagus (fig. 8E,F; table 1) MATERIAL—NMV P157483, posterior part of left P® (fig. 8E); NMV P157481, posterior portion of right P® (fig. 8F); NMv P160154, anterior part of right P?. DESCRIPTION— The P? fragments NMv P157483 and NMV P157481 possess a broad bladelike pos- terolingual cusp and a broad lingual cingulum. Al- though broken, a piece of a labial cingulum can be seen on NMV P157483. A posterobuccal cusp is present on NMV P157481. NMv P160154 pos- sesses a broad lingual and narrower labial cingu- lum. No distinct anterior cusp is present on the blade as in some species of Dendrolagus (e.g., D. lumholtzi). DIsCUSSION— These specimens have been ten- tatively referred to a species of Dendrolagus, rather than to a species of Sthenurus, Simosthenurus, or Procoptodon, on the basis of the presence of a posterobuccal cusp on the P*, which is separated from the posterior end of the blade much more than is seen in any other genus of macropodid. In addition, they often possess a narrow labial cin- gulum and a broad lingual cingulum. The postero- lingual cusp is broad and bladelike, intersecting the main crest at nearly 90°. These premolar frag- ments differ from those of the sthenurines men- tioned above in possessing a lower lingual cingu- lum and less crenulate enamel. The Hamilton fossils assigned to cf. Dendrola- gus are most similar to Dendrolagus bennettianus among living species in the involvement of the posterior-lingual blade of its P® in the formation of a posterior cingulum and a poorly defined cin- gular basin behind the blade. It is smaller than any living Dendrolagus. The posterior width of P® is 4.2-4.9 mm. The narrowest posterior width for P? known in any living species of Dendrolagus is 8.3 mm fora specimen of D. /umholtzi (Groves, 1982). The P?, which is smaller than that of any named species other than D. bennettianus, is very similar, however, to P® fragment(s) assigned to cf. Den- drolagus from the early Pliocene Bow local fauna (Flannery & Archer, 1984). Thylogale Gray, 1837 Thylogale ignis n. sp. (figs. 9-11, 12A; table 3) HOLOTYPE—NMvV P157480, a barely worn right P? (fig. 9A). — (B), buccal (C), and lingual (D) views at approx. <x 5.9. E, NMv P157483, posterior portion of a left P*? shown at approx. x 4.1. F, NMv P157481, posterior portion of a right P*? shown at approx. 4.1. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 19 REFERRED SPECIMENS—I?—PM 4733, left (pl. XXV C in Turnbull & Lundelius, 1970). I’ —Nnmv P54055, right (fig. 9D); NMv P162890, left; NMV P162891, right; NNV P162892, right; PM 4731, right (pl. XXII A,B in Turnbull & Lundelius, 1970). P?—Nnmv P54078, left (fig. 9C). P?—Nmv P54146, right; NMv P158471, left; NMv P157478, left; Nuv P162893, left; pm 17514, left (fig. 9B). M'!—nmv P54086, right (fig. 9E). M?, M3, or M*—pm 16802, right (fig. 9F); pm 17526, % left upper molar. I,—NMV P26419, left (fig. 1OA); NMv P158455, right; NMVv P160133, right; NMv P160150, right; NMV P160152, left; NMv P160160, right; NMv P160180, left; Nuv P160213, left; Nuv P160665, right; NMV P162894, left; NMv P175472, left; pm 4444, left (figured by Turnbull & Lundelius, 1970, pl. XXV D); Pm 16805, left. P,;—NMvV P54111, left; Nuv P54139, right; NMv P157479, right; NMV P157482, left; Nvv P160132, left; NMV P160224, right (fig. 10B). M,—NMv P54100, right; NMv P158472, right (fig. 10C); NMv P159847, left; Nuv P160188, left; NMV P162895; pm 4430, right; pM 4438, left (fig- ured by Turnbull & Lundelius, 1970, pl. XXVI A-D); PM 17520, right. M,—PM 4564, right (figured by Turnbull & Lun- delius, 1970, pl. XXIII B-E). M,.4—NMV P158432, right (figs. 1OD, 11A). M, or M,—NMv P26420, right; pm 4576, right (figured by Turnbull & Lundelius, 1970, pl. XXIV A-D). TYPE LOCALITY AND AGE—See the Introduction for details. DiAGNosis— Thylogale ignis can be distin- guished from other species of Thylogale in the following ways. Thylogale thetis, T. stigmatica, and T. brunii have a more posteriorly positioned groove on I’, a better developed cingulum on P?, and a markedly broader protolophid on M, than occurs in T. ignis. The molars are broader, the protolo- phid of the M, is slightly wider, and the anterior cingulum is shorter in 7. billardierii than in T. ignis (table 3). T. ignis differs from T. christenseni in being larger and having a lower posterolingual cusp on the P* and in having a proportionately shorter P? and P, relative to the molar series. ETYMOLOGY — /gnis means fire in Latin and is a reference to the vulcanism that resulted in the burial of the Hamilton site beneath a lava flow. DESCRIPTION— I? — In the five known specimens of I of 7. ignis, the groove present on the labial 20 face of the tooth is situated between 1.0 and 1.5 mm from its posterior edge. The tooth (fig. 9D) is almost identical in morphology to that of T. bil- lardierii. P?—The P? of T. ignis (fig. 9C) is shorter than P?; its crest is less prominent and is basically like the P? of T. stigmatica. P*?— This tooth consists of a main blade and a posterolingual cusp (fig. 9B). The posterolingual cusp is much lower than the main blade of the tooth. The main blade consists of anterior and posterior cusps with three smaller cuspules be- tween them. The posteriormost cuspule is indis- tinct and tends to merge with the posterior cusp. A distinct lingual cingulum is absent but a small ridge is present basally on the lingual and buccal sides of the tooth. M'—The single M! in the collection is unworn (fig. 9E). Its parastyle is small and the protoloph is poorly developed. The anterior cingulum is re- stricted to the labial half of the tooth. The midlink, postparacrista, and premetacrista are well devel- oped. The metaloph is strongly developed and a very slight postlink is present on its rear face. The posthyposcrista and postmetacrista form a ‘““U”’- shaped posterior cingulum. This tooth is indistin- guishable in morphology from that of 7. billar- dierii but differs from that of 7. stigmatica, T. brunii, and T. thetis in having the anterior cin- gulum further restricted to the buccal side of the tooth. M?, M3, or M4—This tooth (fig. 9F) has a Thy- logale morphology and is tentatively assigned to T. ignis. Its procingulum is broad and prominent, and the anterior moiety is wider than the posterior. I,—The I, of T. ignis (fig. 10H) is virtually in- distinguishable in size and morphology from that of T. billardierii. The enamel is restricted to the labial side of the tooth except for the flanges of enamel present on the ventral and dorsal margins. P,— The P; is formed ofa simple blade (fig. 10B) that is slightly deflected lingually at its posterior end. The blade of the tooth is formed by anterior and posterior cuspids with three smaller cuspules between them. A small cingulum is present around the base of the tooth. The P, is slightly broader than that of 7. billardierii. Lower Molars—The M, is fully lophodont in form, although the protolophid is much narrower than the hypolophid (fig. 10C). The cristid obliqua and paracristid are well developed, and well-de- fined pre- and postmetacristids are present. No posterior cingulum or ornamentation is present on the rear face of the hypolophid. The anterior cin- FIELDIANA: GEOLOGY i | | ; ‘ ; ; 4mm 3mm Fic. 9. Stereophoto pairs of various teeth assigned to Thylogale ignis. All are shown in crown view and D is also shown in buccal view. A, NMV P157480, the holotype, a nearly unworn right P* shown at approx. x 4.1. B, pm 17514, left P? shown at approx. x 5.7. C, NMv P54078, left P? shown at approx. x 5.7. D, NMv P54055, right I? shown in buccal (left) and crown (right) views at approx. x 6.0. E, NMv P54086, right M' shown at approx. x 3.8. F, pm 16802, right upper molar (M?, M3, or M*) shown at approx. x 3.9. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 21 8mm Fic. 10. Stereophoto pairs showing some of the lower teeth of 7. ignis. The cheek teeth are all shown in crown (occlusal) view. A, NMv P26419, left I, shown in dorsal (left) and ventral (right) views at approx. x 2.3. Insert shows 22 FIELDIANA: GEOLOGY 1cm Fic. 11. Drawings of specimens of 7. ignis (A) and Kurrabi pelchenorum (B), part of the holotype specimen. A, NMV P158432, the right mandibular fragment preserving M,_, is drawn at approx. <6.4. Left to right are lingual, crown, and buccal aspects. B, NMv P158425, the P, from the holotype specimen shown, left to right, in buccal crown, and lingual views at approx. x 3.6. The rest of the type is illustrated in Figure 12. 1cm Ce detail of the forming wear facet at the tip of the tooth. B, NMv P160224, right P,; shown at approx. x 4.3. C, NMV P158472, right M, shown at approx. x 4.4. D, NMv P158432 with M,,, right mandibular fragment shown at approx. x 3.0. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 23 ‘ydAI9 SUI UTYIIM pojdnisuN BuIsq J9}1e] 9Yy) ‘suOTIsOd Jodoid J19y) UI oIe "| pue “| “AjJOLID}Ue SUIORJ IOBJANS [LSN]DOO S}I YIM UIBILW IBJOIATL 9Y} MOOG I[QIPUBU 9Y} UTYIIM oI] 0} UOTISOd [BUIOU s}I JO INO po}eIOI UZ0q sey “W IL (AI “By Ul porensnyyi st fg aU) “€7Tx “xoidde 1e UMOYsS ‘YI99} IBJOW JaIY} YIM sqIpueW Yo] [ered e ‘CTPECId AWN ‘WnsouayIjad i1qosiny JO usutdeds odAjOfOY 9y} JO SBuIMeIG “ZI OL] FIELDIANA: GEOLOGY 24 TABLE 3. Measurements (mm) of three species of Thylogale. Length Anterior width Posterior width xX R SD N xX R SD N xX R SD N T. ignis PP? 6:4 36:2-6:8° 427 4 210 1.8=2:3. .216 4 2.1 1.6-2.6 .41 4 T. billardierii 6.5 6.0-7.4 .46 13 2 19=2.4-" 139% 213 3:4 ~3:0-3.8 -224°- 13 T. christenseni 6:5. -6.3-657 . Val7 4 22 ~2.0=2.3: 126 4 25 2. 6=2.8:.0 G10 4 T. ignis P;.. 6.8 6:5-7:1 28 4 222. 1=2:2: 050 4 3.5 3.2-3.8 .25 4 T. billardierii 5:9, <521=6:6: 41 13 1855 1 6=2:0) = 13 Qo 129=256." 3192" ES T. christenseni 5:4 S31=5:9- 527 9 1.7 1.6-1.8 .09 9 1.7 1.6-1.8 .09 9 T. ignis Mt 5:5 — — 1 3.9 _ _ 1 4.4 _ —_ 1 T. billardierii 4.7 4.5-4.8 .15 5 B3°8y) #374—4.1". 255 5 40 3.8-4.2 .18 5 T. christenseni 4.0 3.9-4.2 .15 3 DET 2: O= 258" 3 2.8 2.8-2.9 .06 3 T. ignis M, 4.8 4.7-4.9 .08 4 2°67 7 2:4=2:8)) . 173 4 3:2> 3.0=3.4. 317 4 T. billardierii 4.0 3.9-4.1 .07 5 234; 2.3=23/° 167 5 2:91 23=20h 208 5 T. christenseni 3:5. 7 -3'4—4;5; 06 4 185 1621.9: +18 4 2:0) -..2.0=251 — 205 4 T. ignis M, 5.0 — — 1 2.9 _ _ 1 3.4 _ — 1 T. billardierii 47° -4:3=5:0) ..18> 610 32 219=3:4: “183710 3:8" 322-338 > G19 oO T. christenseni 3.8.0 326—4.05 52150 £26 PA ae Hie PS mer 23 3.7. 2.4—2.9 16 24 T. ignis M, 5.8 — — 1 3.6 _ - 1 _ _ - = T. billardierii ST oLW=oite “c2ou ali) 3:8° 0 -3:4—42." 272, ld _ _ —_ _ T. christenseni 4.1 3.8-4.4 .18 28 3. O'ane2 S—s702) 1D 26 _ _ _ — T. ignis M, 6.9 _ _ 1 4.3 _ _ 1 4.3 — — 1 T. billardierii 6:15 56-6249 9229: 10) 4S -41055:2" 378 10% \455 2337537 4425 10 T. christenseni 4:6, --4:3=4'3 14s -22 3.4: 3.2=3:8. 19 21 314-7 3B 137855 22021 X = mean, R = range, SD = standard deviation, N = sample size. gulum is extremely narrow and elongate compared to that of other species of Thylogale. One specimen, NMV P158132, has M,_, pre- served in order of position (figs. 10D, 11A). M, differs from M, in being larger, in having the lophids subequal in width, and in having the pre- and postmetacristids reduced in size. The an- terior cingulum is also shorter and broader. M,; differs from M, in being larger and in having the anterior cingulum broader and shorter. M, is larger than M; but otherwise is very sim- ilar. These teeth are indistinguishable from those of T. billardierii apart from being slightly more elongate and narrower (table 3). DiscussiON—Turnbull and Lundelius (1970) previously recognized a species of Thylogale in the Hamilton fauna on the basis of a partial I? and I’, an upper molar, probably M3, M‘*, or M®, and four partial upper molars, a partial lower premolar, three lower incisors, and isolated lower molars. Because of uncertainty regarding the association of isolated teeth, they refrained from naming a new species. The new material justifies the naming of a new species. The I>, P?, and M' show characters that differ from all known species of Thylogale. Al- though only three molars are certainly associated in a jaw fragment, the other teeth appear to rep- FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE resent a single species because of size, relative abundance, and morphology. Ride (1957) notes that T. billardierii differs from all other species of Thylogale in having the labial groove of I° situated farther anteriorly, which is the condition in 7. ignis. This character state is regarded as plesiomorphic within macropodids as it is widespread in other macropodine genera. Sev- eral other primitive character states are seen in 7. billardierii and T. christenseni as well as in the newly described 7. ignis, which are not seen in other species of Thylogale. These characters, such as the form of the M, and the morphology of the M! and P?, appear to point to a dichotomy of advanced and primitive species within Thylogale. The polarity of these features and the relationships of the species of Thylogale are discussed exten- sively in Flannery (1989). Kurrabi Flannery and Archer, 1984 Kurrabi pelchenorum n. sp. (figs. 11B, 12--14; table 4) HOLOTYPE—NMV P158425, partial left dentary containing P, (removed; fig. 11B), hypolophid of M,, complete M,_, (fig. 12). 23 Fic. 13. Stereophoto pairs of four specimens referred to Kurrabi pelchenorum, shown in crown views. A, NMV P160209, left P>. B, NMv P160190, right M'. C, NMv P160494, right M,. D, NMv P158634, left M4. 26 FIELDIANA: GEOLOGY 5mm Fic. 14. Drawings of two teeth of Kurrabi pelchenorum (A, B and F, G) and one of Protemnodon sp. (C-E). A, B, pM 17508, left I,. C-E, pm 16818, right P,. F, G, pm 17507, protolophid, right lower molar, probably M;, M,, or M.. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE Zt TABLE 4. Measurements (mm) of teeth referred to Kurrabi pelchenorum. M, M, M; M, M, NMV P158425 (type) 1G 1 NMV P160494 1 a NMV P158634 | _ NMV P54105 AW — NMV P160368 PW — NMV P160209 1 11.4 NMV P160190 L = length, AW = anterior width, PW = posterior width. REFERRED SPECIMENS—NMV P 160209, left P? (fig. 13A); NMV P160190, right M' (fig. 13B); NMv P54120 and NMv P160179, anterior part of pro- toloph of upper molars; NMv P54192, anterior por- tion of right I,; pM 17508, left I, (figs. 14A,B); NMv P54195, protolophid of left M,; Nvv P160210 and NMV P 160494 (fig. 13C), right M,’s; NMv P158632, protolophid of left M,; NMv P160368, hypolophid of left M,;; NMv P54113, right M,; NMv P158634, left M, (fig. 13D); NMv P54105, protolophid of left M,; NMV P158058, right M,, M,, or M.; PM 17507, right lower M, probably M;, M,, or M; (figs. 14F,G). Type LOCALITY AND AGE. —See the Introduction for details. DiaGnosis— Kurrabi pelchenorum can be dis- tinguished from K. mahoneyi in possessing shorter premolars and lower crowned molars. The P? of K. pelchenorum also differs from that of K. ma- honeyi in possessing a better-developed lingual cingulum and in having an anterior edge that slopes more gently to the crown base. K. pelchenorum differs from K. merriwaensis in possessing lower crowned molars with a better-developed lingual cingulum and a more gently sloping anterior mar- gin. ETyMOLOGY— Named for Mr. and Mrs. George Pelchen, the previous owners of the property con- taining the fossil site, who aided us greatly during our early periods of fieldwork at the Hamilton site. DESCRIPTION— P?— The P® (fig. 13A) is com- 28 posed of a main crest, with anterior and posterior cusps, a lingual cingulum, and a posterolingual cusp. Between the anterior and posterior cusps of the main crest are two well-defined ridgelets and a third, less well-defined one. A posterior fossette and well-developed lingual cingulum are present. Upper Molars— The only known M! is unworn (fig. 13B). The protoloph is almost undeveloped, the paracone and protocone forming distinct cusps. The anterior cingulum is formed of the prepara- crista and is poorly developed, ending lingually anterior to the protocone. The midlink is well de- veloped, and an extremely well-developed post- paracrista is present, forming a longitudinal shear- ing blade. The metaloph is well developed. A postlink is present, situated slightly nearer the metacone than the metaconule on the rear face of the metaloph. The posthypocrista is strongly de- veloped and forms a posterior cingulum with a much more weakly developed postmetacrista. Only two anterior fragments of more posterior upper molars are known. One of them possesses a slight forelink. I,—The I, (figs. 14A,B) is almost identical in morphology to that of K. mahoneyi. The buccal side is extensively enameled, and the dorsal and ventral flanges of enamel are present on the lingual face, reaching approximately one-quarter of the way up the tooth. P,—The P, (fig. 11B) in the holotype has a sim- FIELDIANA: GEOLOGY ple bladelike crest, with a slight posterolingual flexion. The posterior cuspid is the highest point on the crest and the anterior cuspid the lowest. Three ridgelets, decreasing in size anteriorly, are present between the cuspids. The anterior edge of the crest slopes in a similar manner (though not so steeply) as in the P*. Lower Molars—M, is represented by two iso- lated right teeth (fig. 13C) and a protolophid, in addition to the fragmented hypolophid in the ho- lotype. There is no posterior cingulum present. The M, is represented in the holotype by a com- plete, barely worn tooth. The anterior cingulum is low, relatively narrow, and anteroposteriorly short. The paracristid originates at the apex of the pro- toconid, is concave buccally, and terminates in the medial part of the anterior cingulum. The proto- lophid and hypolophid are subequal in width. The cristid obliqua originates just below the apex of the hypoconid and terminates slightly labiad of the center of the rear face of the protolophid. The rear face of the hypolophid is nearly vertically ori- ented and no posterior cingulum or other orna- mentation is present. An unerupted M, is part of the holotype. It differs from M, only in being larg- er. Several isolated M,’s are present; one of them (fig. 13D) compares closely to that of the holotype. Discussion — Kurrabi pelchenorum is closest in morphology to K. merriwaensis from the Bow local fauna. It differs from the latter species primarily in possessing lower crowned molars with weaker links, which may suggest habitat differences. One molar, NMV P158058, a right M;, M,, or M,, may represent an extreme variant of K. pelchenorum or possibly an additional species. It differs from the specimens more certainly assigned to K. pel- chenorum in having the lophids placed closer to- gether and, at least on M,, in having weaker links. It is possible that the hypolophid reported by Turnbull and Lundelius (1970) as belonging to a species of Troposodon (NMvV P26422) also repre- sents the hypolophid of an M, of K. pelchenorum. A dentary fragment (Melbourne University Ge- ology Department No. 2014), initially assigned as a sthenurine kangaroo (Stirton, 1957) but later considered to be a species of either Setonix, Den- drolagus, Dorcopsis, or Dorcopsulus (Tedford, 1966), was reported from the marine sediments of Forsyth’s Bank, Grange Burn, near Hamilton. For- syth’s Bank is approximately | km downstream from the Hamilton site (site 8 on Gill’s map of the area; Gill, 1957, text fig. 3) and represents an earlier time period. [? Miocene or earliest Plio- cene?] This specimen is similar in M, morphology to the holotype of K. pelchenorum (the only com- parable tooth between the specimens) and the oth- er species of the genus. However, it differs in being larger (see table 4), and the alveoli for the roots of P, indicate that it was a much larger tooth abso- lutely and relative to molar size than P, in K. pelchenorum. It differs from the other species of Kurrabi in having lower crowned molars. It prob- ably represents yet another distinct species in this genus, but in any case it is not part of the Hamilton fauna. Protemnodon Owen, 1874 Protemnodon sp. (figs. 14C-E, 15; table 1) MATERIAL—NMV P 160372, a left M'! (figs. 15A— E); PM 4429, left M? (figured by Turnbull & Lun- delius, 1970, pl. XXVIII A-E, as a dP?); NMV P162896, anterior fragment of left P® (figs. 15F-— I); PM 16818, nearly complete left P, (figs. 14C- E). DESCRIPTIONS— The P? fragment contains the anterior cusp and the two anteriormost cuspules on the main blade and the anterior portion of the lingual cingulum. The lingual cingulum terminates lingual to the anterior cusp and is composed of a series of tubercles, which correspond to the lingual ridgelets originating at the cusp and cuspules of the main crest. The M! is barely worn and is low-crowned. The protoloph is poorly developed, as is the midlink. The preparacrista joins a poorly developed ante- rior cingulum, which extends only two-thirds the width of the tooth. A well-developed postpara- crista is present. The metaloph is broader and bet- ter developed than the protoloph. The pre- and postmetacristae are well developed as is the post- hypocrista. A weakly developed postlink is present on the rear face of the metaloph, placed slightly closer to the metacone than the hypocone. The M? has been described in Turnbull and Lundelius (1970), where it was considered to be a dP*. The P,, like that of Kurrabi pelchenorum, has a simple bladelike crest with a slight posterior labial flexure. There are three weak ridges ahead of the posterior cuspid. They extend from the crest to the base and decrease slightly in size posteriorly rather than anteriorly as in K. pelchenorum. The tooth minus its anterior cusp is as long as the complete tooth of K. pelchenorum. DIsCUSSION— The M'! is most similar to that of Protemnodon otibandus among named macropo- FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 29 Fic. 15. Drawings of two specimens referred to Protemnodon sp. A-E, NMv P160372, left M' shown in crown (A), buccal (B), posterior (C), anterior (D), and lingual (E) views. F-I, NMv P162896, left P? shown in crown (F), anterior (G), lingual (H), and buccal (I) views. 30 FIELDIANA: GEOLOGY aca tealle ihrvpi We Sak rs m8 tei ae en le, ie Se eee eee doids. However, it differs from P. otibandus in being proportionately broader and in having the anterior cingulum more restricted lingually. The premolar fragments indicate that a distinct ante- rior cusp is present, as in most species of Protem- nodon, and that the P* possesses a broad lingual cingulum. While probably representing a new spe- cies, more complete material is needed before it can be named. Macropus Shaw, 1790 Macropus (Notamacropus) sp. (fig. 16A; table 1) MATERIAL—NMV P54112, hypolophid ofa right lower molar. DESCRIPTION—The specimen is a completely unworn hypolophid, probably from an unerupted tooth. The occlusal edge is much narrower than the base of the crown. The midlink is well devel- oped, originating at the apex of the hypolophid. The accessory link is a sharp raised crest that ex- tends anterolingually from a point just lingual to the cristid obliqua to the point where it is broken away. The hypoconid is taller than the entoconid. DIsCUSSION—The specimen can be recognized as representing a species of Macropus because it possesses a strongly posteriorly convex hypolo- phid. The posterior face of the hypolophid is unor- namented except for an extremely shallow groove. This differs from the condition in species of Mac- ropus (Macropus), where a deeper groove or fos- sette is invariably present. An accessory link buc- cal to the midlink is present in the Hamilton tooth. This has not been observed in species of Macropus (Osphranter) or M. (Macropus) but occurs occa- sionally in species of M. (Notamacropus) (see M. (N.) agilis siva in Bartholomai, 1975). Its small size is also suggestive of a species of Macropus (Notamacropus) (see Dawson & Flannery, 1985). This tooth fragment is the only evidence of a truly hypsodont macropodid in the Hamilton fau- na. Although it is determinable to the subgeneric level, it cannot be assigned to a species. Wallabia Trouessart, 1905 cf. Wallabia (fig. 16B; table 1) MATERIAL—NMV P54208, a single unworn left Be DESCRIPTION—NMV P54208 consists of a blade and a posterolingual cusp. Anterior and posterior cusps are present on the blade, with three distinct cuspules and their ridgelets and a weaker fourth cuspule between them. The posterolingual cusp is connected to the main crest by two ridges, one that extends around the posterior end of the tooth, the other that goes directly to the main crest, inter- secting it at 90°. A broad, well-developed lingual cingulum is present and a weak buccal one. This tooth appears to be indistinguishable in size and morphology from that of Wallabia bicolor. DISCUSSION— This specimen can be distin- guished from the P® of all potoroids and macro- podine species of the genera Dorcopsulus, Dorcop- sis, Lagorchestes, Onychogale, and Macropus and the species of Thylogale billardierii, T. ignis, and T. christenseni by possessing a well-developed lin- gual cingulum. It lacks the distinct lingual crest seen on the P? of sthenurine kangaroos. The tooth is smaller than that of species of Protemnodon. The main blade is more distinctly ridged than that of species of Macropus (Osphranter) and lacks the posterobuccal cusp often seen in species of Den- drolagus. Although possibly belonging to a form close to Setonix, Thylogale, Prionotemnus, Mac- ropus (Notamacropus), or Petrogale, it is here ten- tatively assigned to Wallabia, because it is most similar in morphology to the P?* of W. bicolor. The doubt surrounding the taxonomic status of this specimen, and the poorly understood fossil record of the species of Wallabia, makes further analysis pointless. Macropodinae gen. indet. 1 An isolated premolar probably represents a P, or P, of a macropodine (NMV P158431). It consists of an enlarged posterior cuspid that stands slightly above the rest of the tooth, an anterior cuspid, and three cuspules in between. This tooth resembles the P, of Lagorchestes conspicillatus except that it possesses an extra cuspule. Macropodinae gen. indet. 2 The enamel cap ofa large right I, (NMv P160165; fig. 16C) may representa species near Dorcopsulus. It is too large to belong to D. wintercookorum. It has a slight flange of enamel ventrolingually and is elongate, curving upward anteriorly. FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 31 am aN H . ; + a * 4mm Fic. 16. Stereophoto pairs shown in crown view. A, Macropus (Notamacropus) sp., NMV P54112, hypolophid of a right lower molar. B, Wallabia sp., Nuv P54208, left P?. C, Macropodidae gen. indet. 2, Nuv P160165, right I,. 32 FIELDIANA: GEOLOGY Discussion Four of the Pliocene fossil localities of Austra- lia-New Guinea have been radiometrically dated. These are Hamilton, southwestern Victoria (4.46 + .1 Ma); Bluff Downs, northeastern Queensland (dated to at least 4.0 + .12 Ma; Archer & Wade, 1976); Dog Rocks, southcentral Victoria (>2.1 Ma; Rich et al., 1982); and Awe, Papua New Guinea, with four dates that indicate an age of between 4.5 and 7.6 Ma (Plane, 1967; Hoch & Holm, 1986). However, Page and McDougall (1972) indicate that the dates for this site older than 3.1 Ma may have resulted from incomplete degassing of radiogenic argon at the time of eruption. These assemblages make it possible to examine the biotic and climatic change during the Pliocene. Any attempt to use the habitat preferences and requirements of the living relatives of the Ham- ilton fauna to reconstruct the paleonenvironment faces the possibility that there has been evolu- tionary change in the tolerances of the animals. The first report on the Hamilton fauna (Turnbull & Lundelius, 1970) interpreted the fauna as a rain forest assemblage. The recovery of more material requires that this conclusion be reexamined. Mod- ern representatives of many of the Hamilton taxa are rain forest inhabitants, although they do not all live in the same kind of rain forest. The genera Hypsiprymnodon, Dorcopsis, Den- drolagus, and Strigocuscus have modern represen- tatives that are almost wholly rain forest dwellers. However, the various living species of Dorcopsis live in high mountain forests, lowland rain forest, mossy montane forests, and midmontane forests. Living species of Dendrolagus live mainly in mon- tane rain forests, and Hypsiprymnodon is confined to rain forest, preferring the wetter areas (Johnson & Strahan, 1982). The modern New Guinea spe- cies Strigocuscus gymnotus is primarily a rain for- est animal but has been found in areas of regrowth, mangrove swamps, and woodland savannah (Menzies & Pernetta, 1986). The living species of Thylogale live in diverse environments. T. thetis and T. stigmatica are found in rain forests and wet sclerophyll forests (Ride, 1970). Other species of Thylogale live in high montane forests (7. brunii) or in a variety of hab- itats (T. billardierii). Pseudocheirids are also found in a wide variety of habitats, but, as pointed out in the initial report, there are three taxa of this family in the Hamilton fauna, and the only areas that contain as many taxa today are rain forest situations in New Guinea and Cape York Penin- sula. The living petaurids live in treed areas such as tall forest or tall woodland but not rain forest. Only one of the pygmy possums, Cercartetus caudatus, inhabits rain forest. Burramys today inhabits al- pine or subalpine heath (Calaby et al., 1971; Gul- lan & Norris, 1984; Mansergh, 1984). The harsh environment of the living Burramys is in strong contrast to those of all the other Hamilton taxa. Burramys triradiatus from Hamilton differs mor- phologically from later species and probably dif- fered in its physiological characters as well. Wallabia bicolor prefers wet or dry sclerophyll forests but is found in rain forests, woodland, and heath. The two macropodids with no close living relatives, Protemnodon and Sthenurus, have Pleis- tocene relatives distributed over much of Austra- lia in association with taxa that are not rain forest forms. The various living species of Macropus in- habit a wide variety of habitats but are not rain forest forms. This is the earliest record of Macro- pus, which suggests the presence of open, perhaps grassy areas. A consideration of the entire Hamilton mam- malian assemblage suggests the presence of very diverse habitats in the area. If the living represen- tatives of the Hamilton taxa have not undergone drastic changes in their habitat requirements, the environment must have been much more varied than is seen anywhere today. The Hamilton fauna is one of only two known faunas to have both Trichosurus and Strigocuscus, whose ranges do not overlap today (Flannery et al., 1987). This is an example of a disharmonious association similar to those found in Pleistocene assemblages in Aus- tralia and other parts of the world (Lundelius, 1983). The environment appears to have been more of a mosaic than is seen today, with patches of forest, both rain forest and other wet forests, and open areas. A limited palynological and macroplant analysis has been carried out for Hamilton. Charred stumps of the gymnosperm Phyllocladus are present in growth positions directly under the basalt (figured by Turnbull & Lundelius, 1970, p. 9). In addition, the pollen of Podocarpus, Dacrydium, Phyllocla- dus, Casuarina, Halophagus, Nothofagus, Mur- taceae, and Poacaea have been found (Ladd, pers. comm.). Species of Phyllocladus and Dacrydium are no longer found on mainland Australia. These data further indicate that some rain forest patches were present at the site during the early Pliocene. Today the Hamilton area, where not cleared, sup- ports a Eucalyptus camaldulensis woodland. Palynological studies in the Murray Basin by Martin (1991) indicate a drying trend from the FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 33 mid-Miocene (about 12 million years ago) to the early Pliocene (about 6 million years ago). A brief return to more humid conditions is indicated by the reappearance of gymnosperm and Nothofagus pollen, which is interpreted as rain forest, until just after 5 million years ago. This is very close to the age of the Hamilton fauna. The variable per- centages of Nothofagus and gymnosperm pollen from various localities is interpreted as an indi- cation of a rain forest mosaic. It appears from the existence of the diverse mammalian rain forest assemblages at Hamilton 4.46 million years ago that the restricted modern Australia-New Guinea mammalian rain forest fauna is a relict of a formerly much more wide- spread biota. Faunas such as Bow and Bluff downs from eastern and northern Australia, which rep- resent woodland or riparian habitats, suggest that the rain forest that is thought to have been con- tinuous along the east coast in earlier Tertiary time had begun to fragment by Early Pliocene. Rain forest isolates may have been in existence long enough for considerable endemism to have de- veloped by the Early Pliocene. The small remain- ing areas of southern Australian rain forests today have no unique mammals; however, the Hamilton fauna includes some species such as Milliyowi bun- ganditj, which are not known elsewhere and may represent southern rain forest endemics. The genus Burramys, which is now restricted to montane heaths at higher elevations, seems to have under- gone some evolutionary change since the late Plio- cene (Turnbull et al., 1987). Other Hamilton species may provide clues to the origins of extant forms. The presence of Thy- logale ignis at Hamilton suggests that the ancestors of T. billardierii were rain forest dwellers that sur- vived shrinkage of their habitat by adapting to life in the surviving wet sclerophyll areas. Macropus (Notamacropus) sp. May represent yet another such species, as M. (N.) parma, like T. billardierii, is restricted to southeastern Australia, and both liv- ing species are found in rain forests and wet scle- rophyll forests. Conclusions 1. The Hamilton local fauna provides the ear- liest record for species of the genera Simosthen- urus, Troposodon, Milliyowi, Propleopus, Kurrabi, Thylogale, and Macropus (Notamacropus), as well as possibly Dendrolagus and Wallabia. 34 2. The Hamilton local fauna contains taxa rep- resenting diverse habitats. The fauna may contain elements restricted to the Pliocene southern rain forests. 3. The macropodids from Hamilton show that the macropodoid assemblage typical of the more mesic modern environments of Australia-New Guinea was already present by the Early Pliocene. Acknowledgments Collection of the bulk of the material described here was made possible by grant E7615122 from the Australian Grants Research Committee to Rich. Other material was collected by Turnbull and Lundelius, who received National Science Foundation grant GB 3729. Support also came from the Geology Foundation of the University of Texas and from the Field Museum of Natural History. We would also like to thank Ms. J. Dixon, Curator of Mammals at the Museum of Victoria, Ms. L. Gibson, Collection Manager in Mammals at the Australian Museum, Dr. R. Molnar, Curator of Mammals at the Queensland Museum, and Dr. Bruce Patterson, Curator of Mammals at the Field Museum, for loans of specimens that made this study possible. Mr. and Mrs. George Pelchen and subsequently Dr. and Mrs. Wallace McVicker kindly allowed us access to the site. Mr. and Mrs. Samuel P. Wintercook generously provided their shearer’s quarters for the housing of large field parties in the 1970s and 1980s. Mr. John Calaby gave advice on the specific names. Dr. Tibor Pe- renyi, former staff artist at the Field Museum, and Marlene Werner and Zbignew Jastrzebski, present staff artists, all contributed to the illustrations, as did Ron Testa, former Head of Photography. SEM photographs were made at the Field Museum by Turnbull and R. Wibel. Literature Cited ARCHER, M. 1978. The nature of the molar—premolar boundary in marsupials and a reinterpretation of the homology of marsupial cheekteeth. Memoirs of the Queensland Museum, 18: 157-164. ARCHER, M., AND T. F. FLANNERY. 1985. Revision of the extinct gigantic rat kangaroos (Potoroidae: Mar- supialia), with a description of a new Miocene genus and species and a new Pleistocene species of Propleo- pus. Journal of Paleontology, 59(6): 1331-1349. FIELDIANA: GEOLOGY ARCHER, M., AND M. WADE. 1976. Results of the Ray. E. Lemley Expeditions, Part 1. The Allingham for- mation, a new Pliocene vertebrate fauna from north- ern Queensland. Memoirs of the Queensland Muse- um, 15: 379-397. BARTHOLOMAI, A. 1963. Revision of the extinct mac- ropodid genus Sthenurus Owen in Queensland. Mem- oirs of the Queensland Museum, 15: 21-33. 1967. Troposodon, a new genus of fossil Mac- ropodinae (Marsupialia). Memoirs of the Queensland Museum, 15: 21-33. 1973. The genus Protemnodon Owen (Mar- supialia: Macropodidae) in the upper Cainozoic de- posits of Queensland. Memoirs of the Queensland Mu- seum, 16: 309-363. . 1975. The genus Macropus Shaw (Marsupialia: Macropodidae) in the upper Cainozoic deposits of Queensland. Memoirs of the Queensland Museum, 17: 195-235. 1976. The genus Wallabia Trouessart (Mar- supialia: Macropodidae) in the upper Cainozoic de- posits of Queensland. Memoirs of the Queensland Mu- seum, 17: 375-377. . 1978a. The fossil kangaroos. Australian Mam- malogy, 2: 15-22. . 1978b. The Macropodidae (Marsupialia) from the Allingham formation, Northern Queensland. Re- sults of the Ray. E. Lemley Expeditions, Part 2. Mem- oirs of the Queensland Museum, 18: 127-143. CALABY, J., H. DIMPEL, AND I. M. CowANn. 1971. The mountain pigmy possum Burramys parvus Broom (Marsupialia) in the Kosciusko National Park, New South Wales. CSIRO Division of Wildlife Research Technical Paper 23. CoL.LeTT, R. 1877. Ona collection of mammals from central and northern Queensland. Zoologisches Jahr- buch, 2: 829-940. Dawson, L., AND T. F. FLANNERY. 1985. Taxonomic and phylogenetic status of living and fossil kangaroos and wallabies of the genus Macropus Shaw (Macro- pododiae: Marsupialia) with a new subgeneric name for the large wallabies. Australian Journal of Zoology, 33: 473-498. FLANNERY, T. F. 1983. Revision in the macropodid subfamily Sthenurinae (Marsupialia: Macropodoidea) and the relationships of the species of Troposodon and Lagostrophus. Australian Mammalogy, 6: 15-28. 1989. Phylogeny of the Macropododoidea: A study in convergence, pp. 1-46. Jn Grigg, G., P. Jar- man, and I. Hume, eds., Kangaroos, Wallabies and Rat-Kangaroos. Surrey Beatty & Sons Pty. Ltd., New South Wales. FLANNERY, T. F., AND M. ARCHER. 1984. The Macro- podoidea (Marsupialia) of the early Pliocene Bow local fauna, central eastern New South Wales. Australian Zoologist, 21(4): 357-384. FLANNERY, T. F., M. ARCHER, AND M. PLANE. 1982. Miocene kangaroos (Macropodidae: Marsupialia) from three localities in northern Australia, with a descrip- tion of two new subfamilies. Bulletin of the Bureau of Mineral Resources, Australia, Geology, Geophysics, 7: 287-302. FLANNERY, T. F., W. D. TURNBULL, T. H. V. RICH, AND E. L. LUNDELIuS. 1987. The phalangerids of the early Pliocene Hamilton local fauna, southwestern Victoria, pp. 537-546. Jn Archer, M., ed., Possums and Opos- sums: Studies in Evolution. Surrey Beatty and Sons and the Royal Society of New South Wales, Sydney. GiLt, E. 1952. Range in time of the Australian Tertiary flora. Australian Journal of Science, 15: 47-49. 1957. The stratigraphical occurrence and palaeoecology of some Australian Tertiary marsupials. Memoirs National Museum Victoria, 21: 135-203. GLAUERT, L. 1926. A list of Western Australian fossils. Bulletin of the Geological Survey of Western Austra- lia, 88: 36-71. Gray, J. E. 1821. On the natural arrangement of ver- tebrate animals. London Medical Repository, 15: 296- 310. . 1837. Description of some new or little known Mammalia, principally in the British Museum Col- lection. Magazine Natural History I (Charlesworth), 11: 577-587. Groves, C. P. 1982. The systematics of tree-kangaroos (Dendrolagus; Marsupialia, Macropodidae). Austra- lian Mammalogy, 5: 157-186. GULLAN, P., AND K. Norris. 1984. The habitat of Bur- ramys parvus (Broom) in Victoria, pp. 417-421. In Smith, A. P., and I. D. Hume, eds., Possums and Gliders. Australian Mammal Society, Sydney. Hocn, E., AND P. M. Hom. 1986. New K/Ar deter- minations of the Awe fauna gangue, Papua New Guin- ea: Consequences for late Cenozoic biostratigraphy. Modern Geology, 10: 181-196. JOHNSON, P. M., AND R. STRAHAN. 1982. A further description of the musky rat-kangaroo, Hypsiprym- nodon moschatus Ramsay, 1876 (Marsupialia, Poto- roidae), with notes on its biology. Australian Zoology, 21: 27-46. LONGMAN, H. A. 1924. Some Queensland fossil ver- tebrates. Memoirs of the Queensland Museum 8: 16- 28. LuNDELIusS, E. L., Jk. 1983. Climatic implications of late Pleistocene and Holocene faunal associations in Australia. Alcheringa, 7(2): 125-149. MANSERGH, I. 1984. The mountain pigmy possum (Burramys parvus), (Broom): A review, pp. 413-416. In Smith, A. P., and I. D. Hume, eds., Possums and Gliders. Australian Mammal Society, Sydney. Martin, H.A. 1991. Tertiary stratigraphic palynology and palaeoclimate of the inland river systems in New South Wales, pp. 181-194. Jn Williams, M. A. G., P. De Deckker, and A. P. Kershaw, eds., The Cainozoic of Australia: A Re-appraisal of the Evidence. Special Publication No. 18, Geological Society of Australia, Sydney. MatscuiE, P. 1916. Die Verbreitung der Beuteltiere auf Neuguinea. Mitteilungen des Zoologischen Museum Berlin 8: 257-309. Menzies, J. I., AND J.C. PERNETTA. 1986. A taxonomic revision of cuscuses allied to Phalanger orientalis (Marsupialia: Phalangeridae). Journal of Zoology, Se- ries B, 1: 551-618. MULLER, S., AND H. SCHLEGEL. 1839. Verhandelingen over de Natuurlijke Geschiednis der Nederlandsche FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 35 overzeesche bezittingen, ... . Netherlands Govern- ment, Leiden, pp. 1-228, 49 color pls. OweEN, R. 1874. On the fossil mammals of Australia. Part VIII. Family Macropodidae: Genera Macropus, Osphranter, Phascolagus, Sthenurus, and Protemno- don. Philosophical Transactions of the Royal Society, 164: 245-287, pls. 20-27. PaGE, R. W., ANDI. MCDOUGALL. 1972. Ages of min- eralization of gold and porphyry copper deposits in the New Guinea highlands. Economic Geology, 67: 1034-1048. PARKER, K. L. 1953. Australian Legendary Tales. An- gus and Robertson, Melbourne, 235 pp. PLANE, M. A. 1967. Stratigraphy and vertebrate fauna of the Otibanda formation, New Guinea. Bulletin of the Bureau of Mineral Resources of Australia, Geol- ogy, 86: 1-64. RamsAy, E. P. 1876. Description of a new genus and species of rat-kangaroo, allied to the genus Hypsi- prymnus, proposed to be called Hypsiprymnodon mos- chatus. Proceedings of the Linnean Society of New South Wales, 1: 33-35. Rich, T. H., M. ARCHER, M. D. PLANE, T. F. FLANNERY, N. S. PLEDGE, S. HAND, AND P. V. RicH. 1982. Aus- tralian Tertiary mammal localities, pp. 525-572. In Rich, P. V., and E. M. Thompson, eds., The Fossil Vertebrate Record of Australasia. Monash University Offset Printing Unit, Clayton, Victoria. Ripe, W. D. L. 1957. Protemnodon parma (Water- house) and the classification of related wallabies (Pro- temnodon, Thylogale and Setonix). Proceedings of the Zoological Society of London, 128(3): 327-346. 1961. The cheek teeth of Hypsiprymnodon moschatus Ramsay, 1878. Journal of the Royal So- ciety of Western Australia, 44(2): 367-372. . 1964. A review of Australian fossil marsupials. Journal of the Royal Society of Western Australia, 47: 97-131. . 1970. A Guide to the Native Mammals of Australia. Oxford University Press, London, 249 pp. SCHLEGEL, H., AND S. MULLER. 1845. Over Drie Bui- deltieren uit de Familie der Kengoeroe’s, door Herm. Schlegel en Sal. Miller. 7n Temminck, C. J., ed., Ver- handelingen over de Natuurlijke Gescheidnes der Nederlandsche overseesche bezittingen door de Leden der Natuurkundige Commissie in Indié en andere Schrijvers. Zoologie Leiden, pp. 129-148, pls. 19-24. SHAW, G. 1790. The Naturalist’s Miscellany: Or Col- oured Figures of Natural Objects; Drawn and De- scribed Immediately from Nature, vol. 1. G. Kearsley, London. SKILBECK, C.G. 1980. A preliminary report on the late Cainozoic geology and fossil fauna of Bow, New South Wales. Proceedings of the Linnean Society of New South Wales, 104: 171-180. STEIGER, R. H., AND E. JAGER. 1977. Subcommission on geochronology: Convention on the use of decay constants in geo- and cosmochronology. Earth and Planetary Science Letters, 36: 359-362. STIRTON, R. A. 1957. Tertiary marsupials from Vic- toria, Australia. Memoirs of the National Museum of Victoria, 21: 121-135. 36 Tate, G. H. H. 1948. Results of the Archbold Expe- ditions. No. 59. Studies on the anatomy and phylogeny of the Macropodidae (Marsupialia). Bulletin of the American Museum of Natural History, 91: 239-351. TEDFORD, R. H. 1966. A review of the macropodid genus Sthenurus. University of California Publica- tions, Geological Science, 57: 1-72. TEDFORD, R. H., AND M. O. WooDBURNE. 1987. The Ilariidae, a new family of marsupials from Miocene strata of South Australia and an evaluation of the ho- mology of molar cusps in the Diprotodontia, pp. 401- 418. In Archer, M., ed., Possums and Opossums: Studies in Evolution. Surrey Beatty and Sons and the Royal Society of New South Wales, Sydney. TROUESSART, E.-L. 1905. Catalogus Mammalium tam Viventium quam Fossilium. Quinquennale supple- mentum (1899-1904), Fasc. IV. Cetacea, Edentata, Marsupialia, Allotheria, Monotremata. R. Friedlander und Sohn, Berolini, pp. 753-873. . 1897-1905. Catalogus Mammalium tam Viv- entium quam Fossilium. Nova editio (prima comple- ta). R. Friedlander und Sohn, Berlin, pt. 1 (1897), pp. 1-218; pt. 2 (1897), pp. 219-452; pt. 3 (1897), pp. 453-664; pt. 4 (1898), pp. 665-998; pt. 5 (1898), pp. 999-1264; pt. 6 (1899), pp. 1265-1469; supplement, pts. 1-2 (1904), pp. 1-546; pts. 3-4 (1905), pp. 547- 929. TURNBULL, W. D., ANDE. L. LUNDELIUS, JR. 1970. The Hamilton fauna, a late Pliocene mammalian fauna from the Grange Burn, Victoria, Australia. Fieldiana: Geology, 19: 1-163. TURNBULL, W. D., E. L. LUNDELIUS, JR., AND I. Mc- DOoUGALL. 1965. A potassium—argon dated Pliocene marsupial fauna from Victoria, Australia. Nature, 206: 816. TURNBULL, W. D., T. H. V. RIcH, AND E. L. LUNDELIUS, Jr. 1987. Burramyids (Marsupialia: Burramyidae) of the early Pliocene Hamilton fauna, southwestern Victoria, pp. 729-739. In Archer, M., ed., Possums and Opossums: Studies in Evolution. Surrey Beatty and Sons and the Royal Society of New South Wales, Sydney. West, A. 1974. Aborigines of western Victoria, pp. 62-73. In The Natural History of Western Victoria. Australian Institute of Agricultural Science, Mel- bourne. WoobBuRNE, M. O. 1967. The Alcoota fauna, central Australia: An integrated palaeontological and geolog- ical study. Bulletin of the Bureau of Mineral Resources of Australia, Geology 87: 1-87. Appendix Fossil macropodoid specimens from Hamilton in the collections of the Museum of Victoria and the Field Museum that are not discussed in either this text or in Turnbull and Lundelius (1970), be- cause we could not be certain of their generic or FIELDIANA: GEOLOGY specific identity, are listed here, identified as best $4012, 54025, 54030, 54031, 54037, 54041, we-can: 54044, 54054, 54071, 54073, 54092, 54101, 54117, 54118, 54119, 54121, 54127, 54141, Potoroidae: 54145, 54148, 54152, 54155, 54157, 54162, near Hypsiprymnodon. PM 50907, 50935, 50939, 54171, 54174, 54196, 54206, 158072, 50993 158446, 158447, 158448, 158450, 158456, DAO, SIS EERE AT 158457, 158458, 158459, 158460, 158461, Sa ee re 158463, 158464, 158466, 158469, 160129, 160134, 160135, 160136, 160138, 160139, Rach hadiduc MacieRBe aS 160145, 160155, 160163, 160173, 160175, near Dorcopsis. NMv P26418: pM 4431, 4433, 160176, 160183, 160185, 160187, 160189, Ge a ae see ae aaa 160191, 160192, 160195, 160196, 160197, FE TEE TE RSG Seas 160200, 160201, 160206, 160208, 160216, 16835, 16837, 16838, 17509, 17515, 17526, 160369, 160370; pm 16809, 17516, 17519, 17527, 17539, 50768, 50776, 50889, 50942 50764, 50769, 50775, 50777, 50898 Lower molars: NMv P54001, 54026, 54033, 54042, 54043, 54049, 54053, 54056, 54060, 54062, 54067, 54072, 54091, 54096, 54106, 54116, 54135, 54142, 54143, 54156, 54160, 54163, 54173, 54179, 54195, 54198, 54202, near Dendrolagus. PM 16811, 16812 near Thylogale. pm 4432, 4451, 4452, 4483, 4562, 4574, 4586, 16809, 16833, 17513, 17514, 17518, 17520, 50765, 50937, 50945 near Kurrabi. pM 50892, 50894, 50895, 50925, 158019, 158022, 158071, 158434, 158467, 50926 158468, 160131, 160137, 160140, 160141, near Protemnodon. NMV P160166; PM 17507, 160144, 160146, 160148, 160161, 160162, £7508. 17510).27521 160167, 160168, 160169, 160170, 160171, near Macropus. PM 16806, 50940 160174, 160177, 160193, 160198, 160202, near Wallabia. pm 16826, 50772 160203, 160204, 160205, 160212, 160215, 160218, 160220, 160317, 160371, 160429; indet. Macropodid. pM 16812, 16814, 16824, 16825, 16833, Upper incisors: nmv P54022, 54120, 54129, 16834, 17502, 17503, 17512, 17516, 50770, 157471, 158441, 160149, 160151, 160154, 50775, 50936 160156, 160172, 160219, 160222; pm Premolars and fragments: NMV P54028, 50896 54056, 54078, 54150, 54169, 157476, Lower incisors: NMV P54187, 54190, 157472, 158440, 159788, 160142, 160164, 160182, 158445, 160130, 160145, 160148, 160158, 160195, 160367; PM 16815, 16834, 16839, 160160, 160214: pM 16804, 16813, 16821. 17522, 50773, 50774, 50891, 50940, 50941, 17517, 50890 50946, 50947 Upper molars: NMv P54001, 54002, 54003, FLANNERY ET AL.: MACROPODOIDEA (MARSUPIALIA) OF THE EARLY PLIOCENE 37 a, a = a oe os on A Selected Listing of Other Fieldiana: Geology Titles Available Literature of the Receptaculitid Algae: 1805-1980. By Matthew H. Nitecki, Kristine L. Bradof, and Doris V. Nitecki. Fieldiana: Geology, n.s., no. 16, 1987. 215 pages. Publication 1380, $24.00 The Mammalian Fauna of Madura Cave, Western Australia. Part VII: Macropodidae: Sthenurinae, Macropodinae, with a Review of the Marsupial Portion of the Fauna. By Ernest L. Lundelius, Jr., and William D. Turnbull. Fieldiana: Geology, n.s., no. 17, 1989. 71 pages, 21 illus., 21 tables. Publication 1399, $15.00 The Ear Region in Xenarthrans (= Edentata: Mammalia). Part I. Cingulates. By Bryan Patterson, Walter Segall, and William D. Turnbull. Fieldiana: Zoology, n.s., no. 18, 1989. 46 pages, 17 illus. Publication 1405, $13.00 A Preliminary Survey of Fossil Leaves and Well-Preserved Reproductive Structures from the Sentinel Butte Formation (Paleocene) near Almont, North Dakota. By Peter R. Crane, Steven R. Manchester, and David L. Dilcher. Fieldiana: Geology, n.s., no. 20, 1990. 63 pages, 36 illus., 1 table. Publication 1418, $13.00 Protoptychus hatcheri Scott, 1895. The Mammalian Faunas of the Washakie Formation, Eocene Age, of Southern Wyoming. Part II. The Adobetown Member, Middle Division (= Washakie B), Twka/2 (In Part). By William D. Turnbull. Fie/diana: Geology, n.s., no. 21, 1991. 33 pages, 12 illus., 6 tables. Publication 1421, $13.00 A Catalogue of Type Specimens of Fossil Vertebrates in the Field Museum of Natural History. Classes Amphibia, Reptilia, Aves, and Ichnites. By John Clay Bruner. Fieldiana: Geology, n.s., no. 22, 1991. 51 pages, | illus. Publication 1430, $15.00 The Ear Region in Xenarthrans (~ Edentata: Mammalia). Part II. Pilosa (Sloths, Anteaters), Palaean- odonts, and a Miscellany. By Bryan Patterson, Walter Segall, William D. Turnbull, and Timothy J. Gaudin. Fieldiana: Geology, n.s., no. 24, 1992. 79 pages, 24 illus., 1 table. Publication 1438, $20.00 Order by publication number and/or ask for a free copy of our price list. All orders must be prepaid. Illinois residents add current destination tax. 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